When vessel occlusion is used to generate the decerebrate state, the lateral vestibulospinal tract becomes hyperactive because of damage to Purkinje cells in the anterior vermis of the cerebellum, which provide the major inhibitory projection to the lateral vestibular nucleus. This hypertonia is actually not lost after transection of the dorsal roots, which implies that the lateral vestibulospinal tract is acting to a significant extent directly on α motor neurons (either monosynaptically or via interneurons).
Brainstem Control of Posture and Movement
The importance of motor control pathways that originate in the brainstem is evident from observations of the extensor hypertonus and increased phasic stretch reflexes that occur in decerebrate animals. Particular brainstem systems have been identified as influencing posture and locomotion. Brainstem circuits are also critically involved in the control of eye movement; these circuits are discussed in a separate section at the end of the chapter.
Several reflex mechanisms are evoked when the head is moved or the neck is bent. There are three types of postural reflexes: vestibular reflexes, tonic neck reflexes, and righting reflexes. The sensory receptors responsible for these reflexes include the vestibular apparatus (see ), which is stimulated by head movement, and stretch receptors in the neck.
The vestibular reflexes constitute one class of postural reflex. Rotation of the head activates sensory receptors of the semicircular canals (see ). In addition to generating eye movement, the sensory input to the vestibular nuclei results in postural adjustments. Such adjustments are mediated by commands transmitted to the spinal cord through the lateral and medial vestibulospinal tracts and the reticulospinal tracts. The lateral vestibulospinal tract activates extensor muscles that support posture. For instance, if the head is rotated to the left, postural support is increased on the left side. This increased support prevents the person from falling to the left as the head rotation continues. A person who has any disease that eliminates labyrinthine function in the left ear tends to fall to the left. Conversely, a person with a disease that irritates (stimulates) the left labyrinth tends to fall to the right. The medial vestibulospinal tract causes contractions of neck muscles that oppose the induced movement (vestibulocollic reflex).
Tilting the head also changes the linear acceleration on individual hair cells of the otolith organs of the vestibular apparatus. The resulting changes in hair cell activity can produce eye movement and postural adjustment. For example, when a quadruped, such as a cat, tilts the head and body forward (without bending the neck and consequently without evoking the tonic neck reflexes), the result is extension of the forelimbs and flexion of the hind limbs. This vestibular action tends to restore the body toward its original orientation. Conversely, if the quadruped tilts the head and body backward (without bending the neck), the forelimbs flex and the hind limbs extend. Otolithic organs also contribute to the vestibular placing reaction. If an animal, such as a cat, is dropped, stimulation of the utricles leads to extension of the forelimbs in preparation for landing.
The tonic neck reflexes are another type of positional reflex. These reflexes are activated by the muscle spindles found in neck muscles. These muscles contain the largest concentration of muscle spindles of any muscle in the body. If the neck is bent (without tilting of the head), the neck muscle spindles evoke tonic neck reflexes without interference from the vestibular system. When the neck is extended, the forelimbs extend and the hind limbs flex. The opposite effects occur when the neck is flexed. Note that these effects are opposite those evoked by the vestibular system. Furthermore, if the neck is bent to the left, the extensor muscles in the limbs on the left contract more, and the flexor muscles in the limbs on the right side relax.
CHAPTER 9 Organization of Motor Function